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Microbes in snow and ice ecosystems in polar regions contribute substantially to C, N, and P cycling, but few studies have explored microbial activity in seasonal snow. The purpose of this study was to explore the relative importance of snow microbial processing of C, N, and P compounds in atmospheric deposition and litter and detect elemental limitations of snow microbes in Rocky Mountain conifer forests. Enzyme activity in snow was orders of magnitude greater than activity reported for lentic and lotic waters in similar environments. Proportions of C/P‐ and C/N‐acquiring enzymes suggest that snow samples were P limited, or C and P co‐limited, while lentic and lotic waters were more N limited. As such, microbes in seasonal snow may change the composition of nutrients and carbon, but these processes are vulnerable to changes in atmospheric deposition and snow extent and duration, which could affect nutrient processing across large areas. [ABSTRACT FROM AUTHOR]

The plant family Cactaceae is considered among the most threatened groups of organisms on the planet. The threatened status of the cacti family has created a renewed interest in the highly evolved physiological and morphological traits that underpin their persistence in some of the harshest subtropical environments in the Americas. Among the most important anatomical features of cacti is the modification of leaves into spines, and previous work has shown that the stable isotope chemistry of cacti spines records potential variations in stem water balance, stress, and Crassulacean acid metabolism (CAM). We review the opportunities, challenges, and pitfalls in measuring δ 13C, δ 2H, and δ 18O ratios captured in spine tissues that potentially reflect temporal and spatial patterns of stomatal conductance, internal to atmospheric CO2 partial pressures, and subsequent patterns of photosynthetic gas exchange. We then evaluate the challenges in stable isotope analysis in spine tissues related to variation in CAM expression, stem water compartmentalization, and spine whole-tissue composition among other factors. Finally, we describe how the analysis of all three isotopes can be used in combination to provide potentially robust analysis of photosynthetic function in cacti, and other succulent-stemmed taxa across broad spatio-temporal environmental gradients. [ABSTRACT FROM AUTHOR]

Global changes that alter soil water availability may have profound effects on semiarid ecosystems. Although both elevated CO2 (eCO2) and warming can alter water availability, often in opposite ways, few studies have measured their combined influence on the amount, timing, and temporal variability of soil water. Here, we ask how free air CO2 enrichment (to 600 ppmv) and infrared warming (+ 1.5 °C day, + 3 °C night) effects on soil water vary within years and across wet-dry periods in North American mixed-grass prairie. We found that eCO2 and warming interacted to influence soil water and that those interactions varied by season. In the spring, negative effects of warming on soil water largely offset positive effects of eCO2. As the growing season progressed, however, warming reduced soil water primarily (summer) or only (autumn) in plots treated with eCO2. These interactions constrained the combined effect of eCO2 and warming on soil water, which ranged from neutral in spring to positive in autumn. Within seasons, eCO2 increased soil water under drier conditions, and warming decreased soil water under wetter conditions. By increasing soil water under dry conditions, eCO2 also reduced temporal variability in soil water. These temporal patterns explain previously observed plant responses, including reduced leaf area with warming in summer, and delayed senescence with eCO2 plus warming in autumn. They also suggest that eCO2 and warming may favor plant species that grow in autumn, including winter annuals and C3 graminoids, and species able to remain active under the dry conditions moderated by eCO2. [ABSTRACT FROM AUTHOR]

Abstract: Temporal variation in soil nitrogen (N) availability affects growth of grassland communities that differ in their use and reuse of N. In a 7‐year‐long climate change experiment in a semi‐arid grassland, the temporal stability of plant biomass production varied with plant N turnover (reliance on externally acquired N relative to internally recycled N). Species with high N turnover were less stable in time compared to species with low N turnover. In contrast, N turnover at the community level was positively associated with asynchrony in biomass production, which in turn increased community temporal stability. Elevated CO2 and summer irrigation, but not warming, enhanced community N turnover and stability, possibly because treatments promoted greater abundance of species with high N turnover. Our study highlights the importance of plant N turnover for determining the temporal stability of individual species and plant communities affected by climate change. [ABSTRACT FROM AUTHOR]

Grassland soils are significant carbon (C) sinks as more than half of grassland plant biomass is belowground and roots are the main source of soil C. It is uncertain if grassland soils will continue as C sinks in the future because climate change may affect the dynamic, belowground relationships among crown and root biomass, root chemistry and morphology, and root and soil decomposition, all of which influence C sequestration potential. To better understand future belowground C cycling in semiarid grasslands we analyzed three native species ( Bouteloua gracilis, Carex eleocharis, and Pascopyrum smithii ) and mixed-grass community crown and root biomass, root chemistry, morphology, and decomposability, and soil organic carbon (SOC) priming following seven years of simulated climate change at the Prairie Heating and CO2 Enrichment (PHACE) experiment in Wyoming, USA. We found that individual species and the community respond uniquely to the climate change field treatments, indicating that species composition is important when analyzing climate change effects on grassland C cycling. Root biomass in the C3 sedge, C. eleocharis, increased under elevated CO 2 , especially when combined with warming. Decomposition rates of roots from warming plots were higher than those from ambient plots for B. gracilis and P. smithii . Across species, root decomposition rates increased with C and N concentrations. Root morphology was altered as well: B. gracilis root diameter increased under warming, and P. smithii specific root length and surface area increased under elevated CO 2 . P. smthii roots induced short-term, negative SOC priming across all field treatments. Together, our results indicate that grass roots may play a critical role in maintaining soil C stocks in grasslands in the future. [ABSTRACT FROM AUTHOR]

Determining whether the terrestrial biosphere will be a source or sink of carbon (C) under a future climate of elevated CO2 ( eCO2) and warming requires accurate quantification of gross primary production ( GPP), the largest flux of C in the global C cycle. We evaluated 6 years (2007-2012) of flux-derived GPP data from the Prairie Heating and CO2 Enrichment ( PHACE) experiment, situated in a grassland in Wyoming, USA. The GPP data were used to calibrate a light response model whose basic formulation has been successfully used in a variety of ecosystems. The model was extended by modeling maximum photosynthetic rate ( Amax) and light-use efficiency ( Q) as functions of soil water, air temperature, vapor pressure deficit, vegetation greenness, and nitrogen at current and antecedent (past) timescales. The model fits the observed GPP well ( R2 = 0.79), which was confirmed by other model performance checks that compared different variants of the model (e.g. with and without antecedent effects). Stimulation of cumulative 6-year GPP by warming (29%, P = 0.02) and eCO2 (26%, P = 0.07) was primarily driven by enhanced C uptake during spring (129%, P = 0.001) and fall (124%, P = 0.001), respectively, which was consistent across years. Antecedent air temperature (Tairant) and vapor pressure deficit ( VPDant) effects on Amax (over the past 3-4 days and 1-3 days, respectively) were the most significant predictors of temporal variability in GPP among most treatments. The importance of VPDant suggests that atmospheric drought is important for predicting GPP under current and future climate; we highlight the need for experimental studies to identify the mechanisms underlying such antecedent effects. Finally, posterior estimates of cumulative GPP under control and eCO2 treatments were tested as a benchmark against 12 terrestrial biosphere models ( TBMs). The narrow uncertainties of these data-driven GPP estimates suggest that they could be useful semi-independent data streams for validating TBMs. [ABSTRACT FROM AUTHOR]

Vegetation greenness, detected using digital photography, is useful for monitoring phenology of plant growth, carbon uptake and water loss at the ecosystem level. Assessing ecosystem phenology by greenness is especially useful in spatially extensive, water-limited ecosystems such as the grasslands of the western United States, where productivity is moisture dependent and may become increasingly vulnerable to future climate change., We used repeat photography and a novel means of quantifying greenness in digital photographs to assess how the individual and combined effects of warming and elevated CO2 impact ecosystem phenology (greenness and plant cover) in a semi-arid grassland over an 8-year period., Climate variability within and among years was the proximate driver of ecosystem phenology. Individual and combined effects of warming and elevated CO2 were significant at times, but mediated by variation in both intra- and interannual precipitation. Specifically, warming generally enhanced plant cover and greenness early in the growing season but often had a negative effect during the middle of the summer, offsetting the early season positive effects. The individual effects of elevated CO2 on plant cover and greenness were generally neutral., Opposing seasonal variations in the effects of warming and less so elevated CO2 cancelled each other out over an entire growing season, leading to no net effect of treatments on annual accumulation of greenness. The main effect of elevated CO2 dampened quickly, but warming continued to affect plant cover and plot greenness throughout the experiment. The combination of warming and elevated CO2 had a generally positive effect on greenness, especially early in the growing season and in later years of the experiment, enhanced annual greenness accumulation. However, interannual precipitation variation had larger effect on greenness, with two to three times greater greenness in wet years than in dry years., Synthesis. Seasonal variation in timing and amount of precipitation governs grassland phenology, greenness and the potential for carbon uptake. Our results indicate that concurrent changes in precipitation regimes mediate vegetation responses to warming and elevated atmospheric CO2 in semi-arid grasslands. Even small changes in vegetation phenology and greenness in response to warming and rising atmospheric CO2 concentrations, such as those we report here, can have large consequences for the future of grasslands. [ABSTRACT FROM AUTHOR]

Climate controls vegetation distribution across the globe, and some vegetation types are more vulnerable to climate change, whereas others are more resistant. Because resistance and resilience can influence ecosystem stability and determine how communities and ecosystems respond to climate change, we need to evaluate the potential for resistance as we predict future ecosystem function. In a mixed-grass prairie in the northern Great Plains, we used a large field experiment to test the effects of elevated CO2, warming, and summer irrigation on plant community structure and productivity, linking changes in both to stability in plant community composition and biomass production. We show that the independent effects of CO2 and warming on community composition and productivity depend on interannual variation in precipitation and that the effects of elevated CO2 are not limited to water saving because they differ from those of irrigation. We also show that production in this mixed-grass prairie ecosystem is not only relatively resistant to interannual variation in precipitation, but also rendered more stable under elevated CO2 conditions. This increase in production stability is the result of altered community dominance patterns: Community evenness increases as dominant species decrease in biomass under elevated CO2. In many grasslands that serve as rangelands, the economic value of the ecosystem is largely dependent on plant community composition and the relative abundance of key forage species. Thus, our results have implications for how we manage native grasslands in the face of changing climate. [ABSTRACT FROM AUTHOR]

As global changes reorganize plant communities, invasive plants may benefit. We hypothesized that elevated CO2 and warming would strongly influence invasive species success in a semi-arid grassland, as a result of both direct and water-mediated indirect effects., To test this hypothesis, we transplanted the invasive forb Linaria dalmatica into mixed-grass prairie treated with free-air CO2 enrichment and infrared warming, and followed survival, growth, and reproduction over 4 yr. We also measured leaf gas exchange and carbon isotopic composition in L. dalmatica and the dominant native C3 grass Pascopyrum smithii., CO2 enrichment increased L. dalmatica biomass 13-fold, seed production 32-fold, and clonal expansion seven-fold, while warming had little effect on L. dalmatica biomass or reproduction. Elevated CO2 decreased stomatal conductance in P. smithii, contributing to higher soil water, but not in L. dalmatica. Elevated CO2 also strongly increased L. dalmatica photosynthesis (87% versus 23% in P. smithii), as a result of both enhanced carbon supply and increased soil water., More broadly, rapid growth and less conservative water use may allow invasive species to take advantage of both carbon fertilization and water savings under elevated CO2. Water-limited ecosystems may therefore be particularly vulnerable to invasion as CO2 increases. [ABSTRACT FROM AUTHOR]

The flux of carbon dioxide (CO2) between terrestrial ecosystems and the atmosphere may ameliorate or exacerbate climate change, depending on the relative responses of ecosystem photosynthesis and respiration to warming temperatures, rising atmospheric CO2, and altered precipitation. The combined effect of these global change factors is especially uncertain because of their potential for interactions and indirectly mediated conditions such as soil moisture. Here, we present observations of CO2 fluxes from a multi-factor experiment in semi-arid grassland that suggests a potentially strong climate – carbon cycle feedback under combined elevated [CO2] and warming. Elevated [CO2] alone, and in combination with warming, enhanced ecosystem respiration to a greater extent than photosynthesis, resulting in net C loss over four years. The effect of warming was to reduce respiration especially during years of below-average precipitation, by partially offsetting the effect of elevated [CO2] on soil moisture and C cycling. Carbon losses were explained partly by stimulated decomposition of soil organic matter with elevated [CO2]. The climate – carbon cycle feedback observed in this semiarid grassland was mediated by soil water content, which was reduced by warming and increased by elevated [CO2]. Ecosystem models should incorporate direct and indirect effects of climate change on soil water content in order to accurately predict terrestrial feedbacks and long-term storage of C in soil. [ABSTRACT FROM AUTHOR]

Across many dryland regions, historically grass-dominated ecosystems have been encroached upon by woody-plant species. In this paper, we compare ecosystem water and carbon dioxide (CO2) fluxes over a grassland, a grassland–shrubland mosaic, and a fully developed woodland to evaluate potential consequences of woody-plant encroachment on important ecosystem processes. All three sites were located in the riparian corridor of a river in the southwest US. As such, plants in these ecosystems may have access to moisture at the capillary fringe of the near-surface water table. Using fluxes measured by eddy covariance in 2003 we found that ecosystem evapotranspiration (ET) and net ecosystem exchange of carbon dioxide (NEE) increased with increasing woody-plant dominance. Growing season ET totals were 407, 450, and 639 mm in the grassland, shrubland, and woodland, respectively, and in excess of precipitation by 227, 265, and 473 mm. This excess was derived from groundwater, especially during the extremely dry premonsoon period when this was the only source of moisture available to plants. Access to groundwater by the deep-rooted woody plants apparently decouples ecosystem ET from gross ecosystem production (GEP) with respect to precipitation. Compared with grasses, the woody plants were better able to use the stable groundwater source and had an increased net CO2 gain during the dry periods. This enhanced plant activity resulted in substantial accumulation of leaf litter on the soil surface that, during rainy periods, may lead to high microbial respiration rates that offset these photosynthetic fluxes. March–December (primary growing season) totals of NEE were −63, −212, and −233 g C m−2 in the grassland, shrubland, and woodland, respectively. Thus, there was a greater disparity between ecosystem water use and the strength of the CO2 sink as woody plants increased across the encroachment gradient. Despite a higher density of woody plants and a greater plant productivity in the woodland than in the shrubland, the woodland produced a larger respiration response to rainfall that largely offset its higher photosynthetic potential. These data suggest that the capacity for woody plants to exploit water resources in riparian areas results in enhanced carbon sequestration at the expense of increased groundwater use under current climate conditions, but the potential does not scale specifically as a function of woody-plant abundance. These results highlight the important roles of water sources and ecosystem structure on the control of water and carbon balances in dryland areas. [ABSTRACT FROM AUTHOR]